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Department Biochemistry and Physiology of Nutrition, Deutsches Institut für Ernährungsforschung (German Institute of Human Nutrition) (DIfE), 14558 Bergholz-Rehbrücke, Germany
1To whom correspondence should be addressed.
 |
INTRODUCTION |
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 TOP INTRODUCTION Protein intake, lean body...Effects of high dietary-protein...REFERENCES |
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) the recommended
intakes (WHO 1985
), which is currently considered to be
harmless and, according to public opinion, may be even beneficial.
Usually relatively short-term experimental studies on the effects
of high-protein intakes have been performed, and so the
consequences of longer-term or chronic nutrient intake are
difficult to judge. Epidemiologic evidence, on the other hand, which
aims to circumvent this difficulty, can be flawed by various biases,
confounding effects or even limitations of the data which make it
difficult to obtain clear-cut cause-effect relationships
(Taubes 1995
).
It was stated (WHO 1985
) that "there are no functional
indicators that can usefully be applied in experimental situations to
detect protein inadequacy before clinically detectable changes
occur." This statement is equally pertinent for the determination of
the tolerable upper intake level
(TUL)2
particularly, if adequacy of protein intake in various physiological
situations (i.e., maintenance, growth, pregnancy, lactation) is to be
estimated. Moreover, no extensive body of data exists covering the
issue.
However, there are bits and pieces of information suggesting that there
is no benefit from increasing the dietary-protein intake far above
the recommended intake level. Below we summarize the relevant available
literature and attempt, by making prudent assumptions, to estimate a
TUL which is defined as the highest level of daily intake that is
likely to pose no risk of adverse health effects (Institute of Medicine, Food and Nutrition Board 1999
).
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Protein intake, lean body mass and physical performance |
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TOPINTRODUCTIONProtein intake, lean body...Effects of high dietary-protein...REFERENCES |
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). The answer to this
question is important not only for athletes and body builders who tend
to believe that a high-protein intake is crucial for their physical
performance (Linseisen et al. 1993
) but for all those
conditions characterized by a decreasing lean body mass, as seen in the
elderly (Nair 1995
) and various catabolic situations.
An increase of daily protein intake from 1.35 to 2.62 g ·
kg-1 for 1 mon in the habitual diet (170 kJ ·
kg-1 · d-1) while
performing daily weight training did not affect either strength or
muscle mass (Lemon et al. 1992
). Also the response of
protein turnover to exercise (4 h at 40% VO2
max.) was independent of dietary protein intake (0.9 or 2.5 g ·
kg-1 · d-1) from
isoenergetic diets (Carraro et al. 1990
).
High-protein meals did not enhance the stimulation of myofibrillar
synthesis induced by resistance exercise in muscle of elderly men and
women (Welle and Thornton 1998
). Daily physical exercise
of increasing intensity at neutral energy balance (constant protein
intake 0.57 g · kg-1 ·
d-1) maintained body protein by an improvement
of N utilization (Butterfield and Calloway 1984
). This
suggests that physical activity has a positive effect on N retention
even at a low level of protein intake, provided energy balance is
achieved. Millward et al. (1994)
stated that, "training and energy
intake depress protein needs and habitual protein intake elevates
protein needs." In conclusion, the literature provides no evidence
that protein nutriture and physical performance are improved by
high-protein diets, the more so as Garlick et al. (1999)
considered
that there are currently no methods sensitive enough to detect whether
high-protein intake results in a long-term increase in
functional lean tissue.
|
Effects of high dietary-protein intakes during adulthood |
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TOPINTRODUCTIONProtein intake, lean body...Effects of high dietary-protein...REFERENCES |
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). In a study comparing fat intake of
normal weight, moderately and severely obese subjects, there was a
strong correlation of body mass index (BMI) with fat consumption but
also with protein intake (g · d-1)
(Alfieri et al. 1997
). A positive correlation between
BMI and protein intake (in % total energy) is frequently reported;
however, this may be caused by underreporting of nonprotein energy
intake (Voss et al. 1998
). On the other hand, in
nonobese volunteers, intake of low-fat foods which corresponds to
an increase in protein intake by 2 energy % reduced energy intake and
body weight (Weststrate et al. 1998
). Nevertheless, it
would be worthwhile to investigate the relationship between
protein-intake level and the development of obesity.
A short-term moderate increase of protein intake (from 50 to
82 g/d) in healthy male volunteers resulted in an increase of insulin
secretion as estimated by 24-h C-peptide excretion (Remer et al. 1996
) (Table 1
). Also an adequate (0.74 g · kg-1 ·
d-1) compared to a high-protein intake (1.87
g · kg-1 · d-1) was
associated with a higher overall insulin sensitivity and lower hepatic
glucose output (Linn et al. 1996
). Interestingly, it was
suggested that amino acids are the most important source of
gluconeogenesis, and thus amino acid catabolism may have a much greater
effect on energy equilibrium than has been thought previously
(Jungas et al. 1992
, Reeds et al. 1998
).
In a 50- to 75- y-old Caucasian population, a daily increment of
0.1 g protein · kg-1 was associated with
an increased risk for microalbuminuria, which is a predictor of renal
and cardiovascular disease (Hoogeveen et al. 1998
).
A 1 SD-increase of protein intake increased the risk of
diabetes by 38% in an aboriginal community in Canada (Wolever et al. 1997
). That amino acids may contribute to insulin
resistance was concluded from results of a recent study in myotube
cells (Patti et al. 1998
).
|
).
It has been reported that a chronic high-protein intake is
associated with a range of functional and morphological changes such as
increased urinary nitrogen excretion, vasopressin plasma levels,
creatinine clearance, glomerular filtration rate, kidney hypertrophy,
renal hemodynamics and eicosanoid production in renal tubules
(Bankir and Kriz 1995
, Brändle et al. 1996
, Yanagisawa and Wadi 1998
). In addition,
increased risk of renal cell cancer has been linked to high-protein
intake (Chow et al. 1994
) (Table 1)
, while among white
males with indicators of kidney disease an increased relative risk of
total mortality with an additional 15 g of protein per day [1.25;
95% confidence interval (CI) = 1.09, 1.42] was observed
(Dwyer et al. 1994
). Patients with moderate renal
insufficiency benefit from a low-protein diet by slowing the
deterioration of renal functions (Klahr et al. 1994
,
Maroni and Mitch 1997
). In addition, epidemiological
evidence suggests a relationship between high-protein intake and
prostate cancer (Vlajinac et al. 1997
) (Table 1)
.
Further, the risk of calcium oxalate stone formation has been
associated with the intake of protein, although this relationship has
not been found consistently (Curhan et al. 1993
,
Hiatt et al. 1996
,Trinchieri et al. 1991
). Oxalate is generated from glycine (Elder and Wyngaarden 1960
), but it appears that variations in dietary
oxalate as well as absorption and excretion rates and generation of
oxalate from breakdown of ascorbic acid may disguise a relationship
between protein intake and calcium oxalate nephrolithiasis in
epidemiological studies. However, it was shown in a controlled
nutritional study that a high-protein diet (1.8 g · kg
-1 · d-1) did lead to
higher oxalate excretion by female subjects and higher glycolate
(precursor of oxalate) excretion in both sexes compared to a
low-protein diet (0.6 g · kg -1 ·
d-1) (Holmes et al. 1993
).
An association between high-protein intake, renal acid and calcium
excretion has been reported (Ball and Maughan 1997
,
Remer and Manz 1994
, Trilok and Draper 1989
). This may be due to the oxidation of sulfur amino acids
(methionine, cysteine), resulting in a decreased fractional renal
tubular reabsorption of calcium (Houillier et al. 1996
,
Trilok and Draper 1989
, Zemel 1988
).
Diets based mainly on plant proteins apparently do not augment calcium
loss, presumably because of a higher phosphate intake and a lower
intake of sulfur amino acids (Ball and Maughan 1997
,
Zemel 1988
). However, in respect to protein quality,
animal protein is superior to plant protein, maintaining nitrogen
balance and thus reducing protein intake. Dietary phosphate modifies
the calciuretic effect of proteins, because it increases renal tubular
reabsorption of calcium. The protein-induced calciuria has
potential negative effects on bone health (Barzel and Massey 1998
, Feskanich et al. 1996
, Kerstetter et al. 1999
) although this is controversial (Heaney 1998
, Munger et al. 1999
).
A high-protein intake was found to result in a mild metabolic
acidosis (Frassetto et al. 1998
). Again, it appears that
sulfur (amino acid) content correlated with renal net acid excretion
(Frassetto et al. 1998
). Chronic metabolic acidosis
decreases protein synthesis, increases protein breakdown and may induce
a negative nitrogen balance (Ballmer et al. 1995
).
Recently a decrease in thyroid function in metabolic acidosis was
observed which might partly explain the effects on protein turnover
(Brungger et al. 1997
). High- protein intake (>2 g
· kg-1 · d-1) is also accompanied by a
decrease of plasma levels of glutamine, alanine and glycine
(Maher et al. 1984
, Matthews and Campbell 1992
). The decline in glutamine can also be a function of
metabolic acidosis (Welbourne 1980
), which increases
renal glutamine extraction (Welbourne et al. 1986
) and
decreases glutamine utilization by lymphocytes (Wu and Flynn 1995
). Glutamine homeostasis at a high dietary-protein
intake is maintained by a decreased de novo production of glutamine
(Matthews and Campbell 1992
) and an increase of hepatic
glutaminase expression (Curthoys and Watford 1995
).
In catabolic patients, plasma glutamine concentration decreases
(Jackson et al. 1999
, Newsholme and Calder 1997
). Glutamine is an important fuel for the intestine and
rapidly dividing cells, such as lymphocytes (Newsholme and Calder 1997
), and reduction in plasma glutamine was shown to be
linked to loss of CD4 + T cells in response to anaerobic training
(Hack et al. 1997
). Decreased glutamine concentration
impaired the ability of cultured lymphocytes to produce interleukin
(IL)-2 and to proliferate (Yaqoob and Calder 1997
). Further, glutamine plays a role in the
regulation of gluconeogenesis (Perriello et al. 1997
),
in protein homeostasis (Hankard et al. 1996
) and in an
experimental human model of glutamine depletion, whole body protein
synthesis decreased (Darmaun et al. 1998
). Low plasma
glutamine levels due to a high-protein intake may reduce
immunologic competence and impair body protein synthesis, particularly
when subjects are metabolically stressed.
Based on the foregoing, it is not evident that high-protein intakes
confer any advantage in terms of strength or health. Moreover,
high-protein intakes must be considered in relationship to the
possible untoward consequences mentioned. Due to the lack of systematic
data, a specific TUL cannot yet be set for a healthy adult population.
However, it would be prudent not to increase protein intakes above
those consumed habitually by well-nourished populations in the
technically advanced nations (2 g · kg-1 ·
d-1). This is in accordance with the recent
recommendation by the International Dietary Energy Consultative Group
(Durnin et al. 1999
).
|
FOOTNOTES |
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Manuscript received August 20, 1999. Initial review completed September 29, 1999. Revision accepted December 6, 1999.
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